Introduction

This vignette describes basic usage of the package intended to process several large bedgraph files in R. Methrix provides set of function which allows easy importing of various flavors of bedgraphs generated by methylation callers, and many downstream analysis to be performed on large matrices. The example data analysis presented here uses the following publicly available whole genome bisulfite sequencing data from GEO: GSE90553 Ziller MJ, Ortega JA, Quinlan KA, Santos DP et al. Dissecting the Functional Consequences of De Novo DNA Methylation Dynamics in Human Motor Neuron Differentiation and Physiology. Cell Stem Cell 2018 Apr 5;22(4):559-574.e9. PMID: 29551301

#Load library
library(methrix)
#Genome of your preference to work with
if (!requireNamespace("BiocManager", quietly = TRUE))
    install.packages("BiocManager")

library(BiocManager)

if(!requireNamespace("BSgenome.Hsapiens.UCSC.hg19")) {
BiocManager::install("BSgenome.Hsapiens.UCSC.hg19")
}
library(BSgenome.Hsapiens.UCSC.hg19) 

Download the data

The data can be directly downloaded from GEO, using the GEOquery package. The files will be downloaded into separate subfolders in the working directory.

if(!requireNamespace("GEOquery")) {
BiocManager::install("GEOquery")
}
library(GEOquery)  
filePaths <-  getGEOSuppFiles("GSM3039352")
filePaths <-  getGEOSuppFiles("GSM3039353")
filePaths <-  getGEOSuppFiles("GSM3039354")
filePaths <-  getGEOSuppFiles("GSM3039348")
filePaths <-  getGEOSuppFiles("GSM3039349")
filePaths <-  getGEOSuppFiles("GSM3039350")
bed_files <- list.files(
  path = getwd(),
  pattern = "*bed\\.gz$",
  full.names = TRUE, 
  recursive = T
)

Processing

CpG annotation

As a first step, we need a list of CpG sites in the respective genome. The CpG sites are listed using the respective Bsgenome annotation package. The read_bedgraph function is also able to extract CpG sites on it’s own, however, it might be beneficial to do it separately.

hg19_cpgs <- suppressWarnings(methrix::extract_CPGs(ref_genome = "BSgenome.Hsapiens.UCSC.hg19"))
## -Extracting CpGs
## -Done. Extracted 29,891,155 CpGs from 298 contigs.

Sample annotation

An annotation table is also necessary to perform analyses. The data will be added to the methrix object, as a colData slot.

sample_anno <- data.frame(
  row.names = gsub(
    pattern = paste0("GSM[[:digit:]]*_BiSeq_(WT_D[[:digit:]]{1,2}_R[[:digit:]]).bed.gz"),
    replacement = "\\1",
    x = basename(bed_files)
    ),
  Day = gsub(
    pattern = paste0("GSM[[:digit:]]*_BiSeq_WT_(D[[:digit:]]{1,2})_R[[:digit:]].bed.gz"),
    replacement = "\\1",
    x = basename(bed_files)
    ),
  Replicate = gsub(
    pattern = paste0("GSM[[:digit:]]*_BiSeq_WT_D[[:digit:]]{1,2}_(R[[:digit:]]).bed.gz"),
    replacement = "\\1",
    x = basename(bed_files)
    ),
  stringsAsFactors = FALSE
)
knitr::kable(sample_anno)
Day Replicate
WT_D14_R1 D14 R1
WT_D14_R2 D14 R2
WT_D14_R3 D14 R3
WT_D6_R1 D6 R1
WT_D6_R2 D6 R2
WT_D6_R3 D6 R3

Reading bedGraph files

read_bedgraphs function is a versatile bedgraph reader intended to import bedgraph files generated virtually by any sort of methylation calling program. It requires user to provide indices for chromosome names, start position and other required fields. There are also presets available to import bedgraphs from most common programs such as Bismark (.cov format), MethylDackel, and MethylcTools. In this case, there is no need to define e.g. chr_idx start_idx arguments, the function will automatically assign them. To know the exact parameters, it might worth to take a look at one of the files, to see the column order.

res <- fread(bed_files[1])
head(res)
##    #chrom start   end ratio totalC methC strand next Plus totalC methC Minus
## 1:     10 60024 60026 0.500      2     1      +    G    +      2     1     -
## 2:     10 60088 60090 1.000      2     2      +    G    +      2     2     -
## 3:     10 60108 60110 1.000      3     3      +    G    +      3     3     -
## 4:     10 60139 60141 1.000      6     6      +    G    +      6     6     -
## 5:     10 60141 60143 1.000      7     7      +    G    +      7     7     -
## 6:     10 60162 60164 0.923     13    12      +    G    +     13    12     -
##    totalC methC localSeq
## 1:      0     0    ATCGG
## 2:      0     0    TGCGT
## 3:      0     0    ATCGT
## 4:      0     0    GACGC
## 5:      0     0    CGCGC
## 6:      0     0    TTCGG

The read_bedgraphs function adds CpGs missing from the reference set, and creates a methylation/coverage matrices. Once the process is complete - it returns an object of class methrix which in turn inherits SummarizedExperiment class. methrix object contains ‘methylation’ and ‘coverage’ matrices (either in-memory or as on-disk HDF5 arrays) along with pheno-data and other basic info. This object can be passed to all downstream functions for various analysis. For further details on the data structure, see the SummarizedExperiment package.

  • For computers with large memory, increase speed using vect=TRUE. The function will open multiple files, defined by the vect_batch_size argument. Keep in mind, that increasing the batch size will increase the memory need as well.

  • If vect=FALSE, the function will iterate along the list of files, keeping only one open at a time.

A more detailed description of the read_bedgraphs function is here: read_bedgraphs)

meth <- methrix::read_bedgraphs(
  files = bed_files,
  ref_cpgs = hg19_cpgs,
  chr_idx = 1,
  start_idx = 2,
  M_idx = 5,
  U_idx = 6,
  stranded = FALSE,
  zero_based = TRUE, 
  collapse_strands = FALSE, 
  coldata = sample_anno,
  vect = TRUE,
  vect_batch_size = 4
)
## ----------------------------
## -Preset:        Custom
## --Missing beta and coverage info. Estimating them from M and U values
## -CpGs raw:      29,891,155 (total reference CpGs)
## -CpGs retained: 28,217,448(reference CpGs from contigs of interest)
## ----------------------------
## -Batch:         1/2
## -Processing:    GSM3039348_BiSeq_WT_D14_R1.bed.gz
## --CpGs missing: 291,768 (from known reference CpGs)
## -Processing:    GSM3039349_BiSeq_WT_D14_R2.bed.gz
## --CpGs missing: 213,929 (from known reference CpGs)
## -Processing:    GSM3039350_BiSeq_WT_D14_R3.bed.gz
## --CpGs missing: 391,604 (from known reference CpGs)
## -Processing:    GSM3039352_BiSeq_WT_D6_R1.bed.gz
## --CpGs missing: 288,499 (from known reference CpGs)
## -Batch:         2/2
## -Processing:    GSM3039353_BiSeq_WT_D6_R2.bed.gz
## --CpGs missing: 225,954 (from known reference CpGs)
## -Processing:    GSM3039354_BiSeq_WT_D6_R3.bed.gz
## --CpGs missing: 392,670 (from known reference CpGs)
## -Finished in:  00:04:56 elapsed (00:05:11 cpu)

Potential pitfalls: Too many missing CpGs, if not expected, might indicate that something went wrong.

  • An obvious error is using the wrong reference genome.

  • A common mistake is falsely indicating strandedness and zero or one based positioning. The combination of the two might lead to only using C-s from one strand (due to a 1 bp shift), resulting lower coverage for the sites. A nice description of the zero- and one-based coordination system can be found here

Initial QC and summary

Without any further processing, we can create an interactive html report containing basic summary statistics of the methrix object with methrix_report function.

The report can be accessed here: Initial reports

methrix::methrix_report(meth = meth, output_dir = getwd(), prefix = "initial")

In the report, we can check genome-wide and chromosome based statistics on coverage and methylation, in order to identify potential quality issues, for example:

  • samples with too high or too low genome-wide coverage or on a specific chromosome. In this case, there is also a possibility, that the sample is affected by copy number alterations. Both methylation levels and coverage therefore shows higher variation among cancer samples.

  • samples with altered beta level distribution.

  • A “bump” on the beta level density plot at 0.5 indicates the presence of single nucleotide polymorphisms (SNPs) -> see later.

Filtering

Coverage based filtering

To ensure the high quality of our dataset, the sites with very low (untrustworthy methylation level) or high coverage (technical problem) should not be used. We can mask these CpG sites. Please note, that the DSS we were using for Differential Methylation Region (DMR) calling, doesn’t need the removal of the lowly covered sites, because it takes it into account by analysis. However, using a not too restrictive threshold won’t interfere with DMR calling.

meth <- methrix::mask_methrix(meth, low_count = 2, high_quantile = 0.99) 
## -Masking coverage lower than 2
## -Masked 168941 CpGs due to too low coverage in sample WT_D14_R1.
## -Masked 87370 CpGs due to too low coverage in sample WT_D14_R2.
## -Masked 233436 CpGs due to too low coverage in sample WT_D14_R3.
## -Masked 160610 CpGs due to too low coverage in sample WT_D6_R1.
## -Masked 100849 CpGs due to too low coverage in sample WT_D6_R2.
## -Masked 234077 CpGs due to too low coverage in sample WT_D6_R3.
## 
## 
## -Masking coverage higher than 99 percentile
## -Masked 268594 CpGs due to too high coverage in sample WT_D14_R1.
## -Masked 251460 CpGs due to too high coverage in sample WT_D14_R2.
## -Masked 225617 CpGs due to too high coverage in sample WT_D14_R3.
## -Masked 254079 CpGs due to too high coverage in sample WT_D6_R1.
## -Masked 275196 CpGs due to too high coverage in sample WT_D6_R2.
## -Masked 264970 CpGs due to too high coverage in sample WT_D6_R3.
## -Finished in:  11.3s elapsed (8.640s cpu)

Later we can remove those sites that are not covered by any of the samples.

meth <- methrix::remove_uncovered(meth) 
## -Removed 219,937 [0.78%] uncovered loci of 28,217,448 sites
## -Finished in:  3.860s elapsed (2.490s cpu)

We might want to remove the sites that are sparsely covered. The function coverage_filter has multiple options on how to perform this. We can set a threshold on the coverage and use either min_samples (minimum number of samples) or prop_samples (minimum proportion of the samples) to define the minimum size of samples that should excess the threshold for each CpG side. bsseq requires

#meth <- methrix::coverage_filter(m=meth, cov_thr = 5, min_samples = 2)
#or
meth <- methrix::coverage_filter(m=meth, cov_thr = 5, min_samples = 2, group="Day")
## -Retained 26,452,492 of 27,997,511 sites
## -Finished in:  4.110s elapsed (2.960s cpu)

SNP filtering

SNPs, mostly the C > T mutations can disrupt methylation calling. Therefore it is essential to remove CpG sites that overlap with common variants, if we have variation in our study population. For example working with human, unpaired data, e.g. treated vs. untreated groups. During filtering, we can select the population of interest and the minimum minor allele frequency (MAF) of the SNPs. SNP filtering is currently implemented for hg19 and hg38 assemblies.

if(!requireNamespace("GenomicScores")) {
BiocManager::install("GenomicScores")
}
library(GenomicScores) 
meth <- methrix::remove_snps(m = meth, keep = TRUE)
## Used SNP database: MafDb.1Kgenomes.phase3.hs37d5.
## Number of SNPs removed:
##       chr      N
##  1:  chr1 258741
##  2: chr10 168199
##  3: chr11 155313
##  4: chr12 155109
##  5: chr13 104670
##  6: chr14 104554
##  7: chr15  96932
##  8: chr16 113226
##  9: chr17 112999
## 10: chr18  88371
## 11: chr19  99802
## 12:  chr2 259890
## 13: chr20  82509
## 14: chr21  46927
## 15: chr22  61581
## 16:  chr3 206538
## 17:  chr4 203468
## 18:  chr5 188502
## 19:  chr6 195152
## 20:  chr7 190251
## 21:  chr8 166357
## 22:  chr9 136295
## 23:  chrX 102053
## 24:  chrY   2001
##       chr      N
## Sum:
## [1] 3299440
## -Finished in:  00:01:29 elapsed (00:01:15 cpu)
  • There is a possibility to keep the sites possibly overlapping with SNPs. We can use this data for detecting possible sample mismatches. For this, we need to plot the most variable sites (probably the ones that are variable in our study population) and plot them on a heatmap.
if (!requireNamespace("pheatmap", quietly = TRUE))
    install.packages("pheatmap")

snps <- meth[[2]]
meth <- meth[[1]]

pheatmap::pheatmap(get_matrix(order_by_sd(snps)[1:min(5000, length(snps)),]))

  • SNP removal in mouse data: Usually, there is no need to remove SNPs from mouse data, since for many experiments, the same strain of mice is used. However, there are cases, when different crosses of strains are used, for which, the level of heterozygosity might be different. SNP data for mice can be downloaded from here: https://www.sanger.ac.uk/science/data/mouse-genomes-project. The SNPs can be filtered out using the region filter function.

Visualization and QC after filtering

Methrix report

After filtering, it worth running a report again, to see if any of the samples were so lowly covered that it warrants an action (e.g. removal of the sample)

Important: Don’t forget to set the recal_stats to TRUE, since the object changed since reading in. The output directory has to be different from last time, to avoid using the intermediate files calculated during the last run.

methrix::methrix_report(meth = meth, output_dir = getwd(), recal_stats = TRUE, prefix="processed")

The report can be found here: processed_methrix_reports.html

There are additional possibilities to visualize the study. We can look at the density of coverage both sample- and group-wise:

methrix::plot_coverage(m = meth, type = "dens")

methrix::plot_coverage(m = meth, type = "dens", pheno = "Day", perGroup = TRUE)

We can visualize te beta value distribution as a violin plot or density plot. These plots (as well as the coverage plot) use the most variable 25000 CpG sites to ensure fast processing. plot_density and plot_violin has the option to use data only from a restricted region.

methrix::plot_density(m=meth)

methrix::plot_density(m=meth, ranges = GRanges("chr22:48007313-49007313"))

Dimensionality reduction

Methrix offers principal component analysis (PCA) to conduct and visualize dimensionality reduction. As a first step, the PCA model has to be calculated. As for plotting, a given number of sites are selected, either random or based on variance (var option). This ensures that the calculations remain feasible.

mpca <- methrix::methrix_pca(m= meth, top_var = 10000, n_pc = 20)

At visualization, we can provide the methrix object to allow color or shape annotation of groups or samples.

methrix::plot_pca(mpca, m=meth, col_anno = "Day", shape_anno = "Replicate")

methrix offers the possibility of region based filtering. With this option, selected regions, e.g. promoters can be visualized. We will use the annotatr package to assess basic annotation categories.

if(!requireNamespace("annotatr")) {
BiocManager::install("annotatr", update = F)
}
library(annotatr)
promoters = build_annotations(genome = 'hg19', annotations = "hg19_genes_promoters")
promoters
## GRanges object with 82960 ranges and 5 metadata columns:
##                 seqnames        ranges strand |             id       tx_id
##                    <Rle>     <IRanges>  <Rle> |    <character> <character>
##       [1]           chr1   10874-11873      + |     promoter:1  uc001aaa.3
##       [2]           chr1   10874-11873      + |     promoter:2  uc010nxq.1
##       [3]           chr1   10874-11873      + |     promoter:3  uc010nxr.1
##       [4]           chr1   68091-69090      + |     promoter:4  uc001aal.1
##       [5]           chr1 320084-321083      + |     promoter:5  uc001aaq.2
##       ...            ...           ...    ... .            ...         ...
##   [82956] chrUn_gl000237     2687-3686      - | promoter:82956  uc011mgu.1
##   [82957] chrUn_gl000241   36876-37875      - | promoter:82957  uc011mgv.2
##   [82958] chrUn_gl000243   10501-11500      + | promoter:82958  uc011mgw.1
##   [82959] chrUn_gl000243   12608-13607      + | promoter:82959  uc022brq.1
##   [82960] chrUn_gl000247     5817-6816      - | promoter:82960  uc022brr.1
##               gene_id      symbol                 type
##           <character> <character>          <character>
##       [1]   100287102     DDX11L1 hg19_genes_promoters
##       [2]   100287102     DDX11L1 hg19_genes_promoters
##       [3]   100287102     DDX11L1 hg19_genes_promoters
##       [4]       79501       OR4F5 hg19_genes_promoters
##       [5]        <NA>        <NA> hg19_genes_promoters
##       ...         ...         ...                  ...
##   [82956]        <NA>        <NA> hg19_genes_promoters
##   [82957]        <NA>        <NA> hg19_genes_promoters
##   [82958]        <NA>        <NA> hg19_genes_promoters
##   [82959]        <NA>        <NA> hg19_genes_promoters
##   [82960]        <NA>        <NA> hg19_genes_promoters
##   -------
##   seqinfo: 93 sequences (1 circular) from hg19 genome
mpca <- methrix::methrix_pca(m= meth, top_var = 10000, ranges = promoters, n_pc = 3, do_plot = F)
methrix::plot_pca(mpca, m=meth, col_anno = "Day", shape_anno = "Replicate")

Differential methylation calling

DSS

A very important step of a WGBS data analysis is the differential methylation calling. During this step, we would like to identify changes between groups or conditions first on the site level. Then, we will need to identify larger regions (DMR-s, differentially methylated regions) affected by methylation changes that are more likely to represent functional alterations. Methrix doesn’t have a DMR caller, therefore we will use DSS. At this stage, DSS is not yet able to work directly with methrix, therefore we transform our methrix object to bsseq for the sake of DMR calling.

if(!requireNamespace("DSS")) {
BiocManager::install("DSS")
}
library(DSS)

For more detailed description, options and recommendations on smoothing, please refer to the ´DSS´ vignette

bs_obj <- methrix::methrix2bsseq(meth)
dmlTest = DSS::DMLtest(bs_obj, group1=rownames(sample_anno)[sample_anno$Day=="D6"], group2=rownames(sample_anno)[sample_anno$Day=="D14"], 
                       smoothing = TRUE)

Call DMRs and DMLs

dmls = DSS::callDML(dmlTest, p.threshold = 0.05)
dmrs = DSS::callDMR(dmlTest, p.threshold = 0.05)

DMRs:

kable(dmrs) %>%
  kable_styling(bootstrap_options = "striped", full_width = F) %>%
  scroll_box(width = "100%", height = "200px")
chr start end length nCG meanMethy1 meanMethy2 diff.Methy areaStat
18184 chr5 92930350 92933392 3043 126 0.9161132 0.8361892 0.0799240 430.6031757
10721 chr19 1466497 1467764 1268 130 0.8324993 0.7562613 0.0762380 398.1324659
18918 chr5 172110452 172111266 815 124 0.7757884 0.6886458 0.0871426 397.2263077
23288 chr9 67340092 67341092 1001 124 0.7617311 0.6736125 0.0881186 389.9699682
24064 chr9 140917238 140918281 1044 115 0.8808357 0.7970408 0.0837949 387.7212469
18174 chr5 92905455 92907417 1963 105 0.8899426 0.8149610 0.0749816 332.6726535
5690 chr12 119591178 119592343 1166 52 0.9016888 0.7620138 0.1396749 266.7463729
16835 chr4 81123867 81125071 1205 109 0.9572686 0.9195279 0.0377407 257.8624785
9500 chr17 35299966 35300973 1008 88 0.9608659 0.9109546 0.0499113 257.1902067
7418 chr15 23378003 23378477 475 70 0.7396353 0.8312258 -0.0915905 -242.9903165
8828 chr16 67428155 67428884 730 74 0.8173113 0.7313175 0.0859938 238.9208790
17587 chr5 3602296 3603053 758 73 0.9230518 0.8598057 0.0632460 230.3238976
9030 chr16 85482200 85483057 858 39 0.5658756 0.7477734 -0.1818979 -229.2458650
11467 chr19 58545241 58545602 362 52 0.8476294 0.7269847 0.1206447 228.1968781
7485 chr15 32604648 32605552 905 84 0.8735001 0.7957639 0.0777362 221.6193792
24917 chrX 154687619 154688089 471 61 0.8923171 0.9638567 -0.0715396 -217.2915896
11627 chr2 19550542 19552098 1557 80 0.9318175 0.8732531 0.0585644 215.7408752
5873 chr12 133158579 133159775 1197 80 0.8130105 0.7267270 0.0862835 215.1361632
20402 chr7 1686521 1687234 714 36 0.5190999 0.7063281 -0.1872282 -212.7787620
19401 chr6 34202568 34204117 1550 75 0.9514510 0.9004377 0.0510133 209.2591448
8061 chr15 83208636 83209267 632 54 0.9067730 0.9756649 -0.0688919 -208.2984354
19944 chr6 114180939 114181421 483 67 0.7936540 0.7004082 0.0932458 205.0188056
24699 chrX 118407207 118407749 543 61 0.7798505 0.8655921 -0.0857416 -201.9642768
1330 chr1 148901580 148902470 891 82 0.7902229 0.7290555 0.0611674 199.9959941
10722 chr19 1467865 1468596 732 78 0.7855306 0.7156071 0.0699235 197.4236394
14352 chr21 47717388 47717995 608 58 0.7422719 0.6433879 0.0988839 196.2631380
55 chr1 3086204 3086757 554 30 0.5501769 0.7667541 -0.2165772 -195.7194311
8226 chr16 744473 745216 744 51 0.9259504 0.8331650 0.0927854 192.4927404
8633 chr16 51147276 51148589 1314 55 0.8220703 0.9068076 -0.0847373 -191.6223407
23333 chr9 70427828 70428417 590 43 0.8946986 0.7879095 0.1067891 190.6655881
9326 chr17 17712912 17713848 937 76 0.8956306 0.8361255 0.0595051 188.8777179
164 chr1 10732172 10733168 997 44 0.5146199 0.6629956 -0.1483756 -187.6521511
20810 chr7 44364754 44365222 469 76 0.9368358 0.9736645 -0.0368287 -187.4400583
16277 chr4 206339 207049 711 55 0.8929468 0.8131918 0.0797551 186.0109428
21647 chr7 150497245 150497760 516 43 0.8104609 0.9252782 -0.1148172 -183.6674497
18183 chr5 92928896 92930058 1163 35 0.9078527 0.7729696 0.1348831 182.4273477
8340 chr16 11035470 11036019 550 53 0.9111264 0.9653631 -0.0542367 -181.2806190
19027 chr6 291882 292507 626 76 0.7850652 0.7177793 0.0672859 181.2005392
8914 chr16 73583735 73584583 849 22 0.5314322 0.8001870 -0.2687548 -181.1114901
14052 chr20 61992070 61992672 603 64 0.9385330 0.8796125 0.0589205 180.6850021
14734 chr3 2140307 2140849 543 62 0.8310748 0.7541629 0.0769118 180.1301515
3525 chr11 6651396 6651836 441 75 0.9064237 0.8583220 0.0481017 179.7342574
21636 chr7 148959279 148959768 490 72 0.9133846 0.9521503 -0.0387657 -177.9939244
14920 chr3 27674210 27674638 429 59 0.7744789 0.6874541 0.0870247 177.4334167
22283 chr8 61834282 61835406 1125 60 0.9546390 0.9005933 0.0540456 175.5054678
198 chr1 11985793 11986233 441 48 0.9153911 0.8370006 0.0783905 174.9610229
21668 chr7 151078463 151078936 474 57 0.8935924 0.8206556 0.0729367 168.9551994
20087 chr6 137809771 137810609 839 62 0.9586632 0.9147548 0.0439083 164.1546140
20417 chr7 2563611 2564603 993 52 0.9406961 0.8693615 0.0713346 163.6274115
22099 chr8 37005133 37005969 837 18 0.5522395 0.8317034 -0.2794640 -163.0567272
2397 chr10 22766224 22767049 826 68 0.9189721 0.8719932 0.0469788 160.6201279
12334 chr2 114341780 114342297 518 54 0.9378503 0.8863733 0.0514770 159.8105971
5053 chr12 52652109 52652599 491 63 0.8118694 0.7465880 0.0652814 157.5257607
12159 chr2 88124876 88125154 279 55 0.8896277 0.9429125 -0.0532847 -156.4360830
17081 chr4 120221435 120222006 572 42 0.9846283 0.9377532 0.0468751 155.1646231
13546 chr20 18568595 18569053 459 54 0.9296749 0.9719525 -0.0422776 -154.6337100
8207 chr15 102501508 102501930 423 52 0.7904408 0.8603573 -0.0699165 -154.1827531
10909 chr19 12175461 12176056 596 57 0.8914417 0.9424102 -0.0509685 -153.9675334
19171 chr6 18155547 18155957 411 47 0.9084892 0.9700963 -0.0616071 -153.4126349
22886 chr8 146228110 146228704 595 54 0.8399533 0.9040175 -0.0640642 -153.0066481
23251 chr9 45762383 45763118 736 53 0.7973017 0.7068416 0.0904601 152.9774211
17551 chr5 2112260 2113123 864 28 0.5132639 0.6901543 -0.1768904 -152.7971237
17596 chr5 4230473 4231862 1390 45 0.7147000 0.8137415 -0.0990414 -151.3539688
7832 chr15 65688225 65688661 437 52 0.7588646 0.6726624 0.0862022 150.6901577
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2755 chr10 72898318 72898403 86 4 0.5195123 0.6028620 -0.0833497 -8.4202360
17100 chr4 124571217 124571421 205 4 0.9156266 0.9589313 -0.0433047 -8.4179510
11208 chr19 38432640 38432810 171 4 0.5391056 0.6224077 -0.0833021 -8.4158570
19976 chr6 117276230 117276286 57 4 0.6784957 0.7549536 -0.0764579 -8.4049162
20733 chr7 34795777 34796017 241 4 0.6018440 0.6864575 -0.0846136 -8.4028511
4069 chr11 69631928 69631995 68 4 0.8779477 0.8339603 0.0439874 8.4020300
19306 chr6 27529740 27529827 88 4 0.7754921 0.8312483 -0.0557563 -8.4007008
14939 chr3 30273946 30274137 192 4 0.5129168 0.5993766 -0.0864598 -8.3957426
17585 chr5 3513855 3513969 115 4 0.5194488 0.5946360 -0.0751872 -8.3913560
24099 chrX 2417134 2417259 126 4 0.7427049 0.8215203 -0.0788154 -8.3904507
17552 chr5 2149388 2149464 77 4 0.5546661 0.6241420 -0.0694759 -8.3897042
4257 chr11 94487388 94487614 227 4 0.5504064 0.6374764 -0.0870700 -8.3883838
19418 chr6 36294310 36294410 101 4 0.5428206 0.6344407 -0.0916200 -8.3877484
729 chr1 51983650 51983732 83 4 0.9358885 0.8901498 0.0457387 8.3809546
19404 chr6 34707640 34707708 69 4 0.5587533 0.6604710 -0.1017177 -8.3794971
15744 chr3 129024838 129024924 87 4 0.9758834 0.9391684 0.0367151 8.3772601
8845 chr16 68936423 68936556 134 4 0.9142139 0.8498203 0.0643936 8.3750521
19282 chr6 26614895 26614947 53 4 0.8756563 0.9231236 -0.0474673 -8.3735799
22544 chr8 98790836 98791046 211 4 0.7661877 0.8283614 -0.0621738 -8.3727349
2693 chr10 63659842 63660050 209 4 0.8646029 0.7997827 0.0648202 8.3667348
4525 chr11 120080712 120080812 101 4 0.7492199 0.6799740 0.0692459 8.3636860
20581 chr7 23058363 23058468 106 4 0.7507088 0.8269791 -0.0762703 -8.3630799
6584 chr13 112725258 112725321 64 4 0.7568896 0.6801339 0.0767557 8.3627875
10333 chr18 38157638 38157727 90 4 0.5382158 0.6210554 -0.0828396 -8.3557936
14561 chr22 33390207 33390319 113 4 0.7270823 0.8070122 -0.0799299 -8.3548190
11556 chr2 9815801 9815856 56 4 0.6086346 0.6867323 -0.0780978 -8.3499579
5960 chr13 28553124 28553347 224 4 0.9271529 0.9640124 -0.0368595 -8.3498890
21313 chr7 102135563 102135629 67 4 0.8175475 0.7684593 0.0490882 8.3429331
20440 chr7 4832009 4832064 56 4 0.6581375 0.7178058 -0.0596683 -8.3425734
14490 chr22 27748424 27748582 159 4 0.5448704 0.6279134 -0.0830430 -8.3417547
16177 chr3 185216097 185216194 98 4 0.8019022 0.8568679 -0.0549657 -8.3383279
20392 chr7 1278832 1279024 193 4 0.9113686 0.9511266 -0.0397580 -8.3316529
15620 chr3 115412640 115412872 233 4 0.5326925 0.6099096 -0.0772171 -8.3299740
2297 chr10 12110679 12110736 58 4 0.8778514 0.8286671 0.0491843 8.3282565
16996 chr4 106394338 106394391 54 4 0.8793332 0.9220707 -0.0427374 -8.3250231
18203 chr5 95047042 95047210 169 4 0.7422754 0.6765624 0.0657130 8.3204488
7644 chr15 48052516 48052587 72 4 0.5176777 0.6022435 -0.0845659 -8.3198533
21286 chr7 100433865 100433983 119 4 0.8387663 0.7570164 0.0817499 8.3135666
319 chr1 21521789 21521902 114 4 0.8070932 0.8759522 -0.0688591 -8.3129726
9165 chr17 4269637 4269704 68 4 0.9430041 0.9732611 -0.0302570 -8.3074444
9964 chr17 71897854 71897906 53 4 0.8615425 0.8075068 0.0540358 8.3061138
5619 chr12 114773266 114773500 235 4 0.6808138 0.7752880 -0.0944742 -8.3027160
13304 chr2 235325925 235326102 178 4 0.5037492 0.5893282 -0.0855790 -8.2919998
1311 chr1 147790557 147790658 102 4 0.8525217 0.8039298 0.0485919 8.2914876
22557 chr8 101541207 101541270 64 4 0.5083866 0.5926474 -0.0842608 -8.2895367
20556 chr7 20834345 20834411 67 4 0.8484594 0.8991494 -0.0506900 -8.2858392
6908 chr14 56424872 56424943 72 4 0.9319439 0.9700849 -0.0381410 -8.2823809
15239 chr3 62694292 62694529 238 4 0.6544842 0.7421437 -0.0876595 -8.2823736
8624 chr16 50776882 50777169 288 4 0.8424948 0.7678821 0.0746127 8.2812613
14919 chr3 27410883 27410958 76 4 0.9651748 0.9345950 0.0305798 8.2807620
12051 chr2 73339084 73339141 58 4 0.9512569 0.9155598 0.0356971 8.2796247
6783 chr14 34567030 34567108 79 4 0.5335846 0.6153074 -0.0817228 -8.2787784
7101 chr14 77278379 77278473 95 4 0.8668063 0.8153137 0.0514926 8.2764335
6161 chr13 51746339 51746468 130 4 0.8270150 0.7673503 0.0596647 8.2739846
5155 chr12 57918829 57918927 99 4 0.5186474 0.6126526 -0.0940053 -8.2738406
14426 chr22 21610190 21610283 94 4 0.6362075 0.7119853 -0.0757778 -8.2715608
2950 chr10 88625155 88625253 99 4 0.7750278 0.6880479 0.0869799 8.2689329
10174 chr18 9102946 9103015 70 4 0.9636644 0.9340244 0.0296400 8.2679330
7683 chr15 53080253 53080315 63 4 0.9701023 0.9393437 0.0307587 8.2658945
4551 chr11 122312560 122312776 217 4 0.8029495 0.7214709 0.0814786 8.2620124
11262 chr19 42667477 42667654 178 4 0.5268543 0.6084816 -0.0816273 -8.2608848
24469 chrX 68484541 68484663 123 4 0.8394741 0.7617561 0.0777179 8.2597710
13688 chr20 34959729 34959921 193 4 0.6116604 0.7087301 -0.0970697 -8.2542762
14739 chr3 2519387 2519503 117 4 0.5269346 0.6166697 -0.0897351 -8.2530089
18151 chr5 92031403 92031543 141 4 0.5191191 0.6070360 -0.0879169 -8.2448855
20449 chr7 5458272 5458356 85 4 0.8890508 0.9326142 -0.0435634 -8.2413494
5999 chr13 31603906 31604100 195 4 0.5168304 0.6060850 -0.0892546 -8.2408804
20064 chr6 134237901 134237970 70 4 0.8846985 0.9304774 -0.0457789 -8.2397786
11025 chr19 20278756 20278819 64 4 0.7450189 0.8011035 -0.0560846 -8.2382147
8908 chr16 73513808 73513913 106 4 0.5135917 0.6051693 -0.0915777 -8.2379583
1313 chr1 147874427 147874498 72 4 0.9167092 0.9524338 -0.0357247 -8.2364233
22772 chr8 135807458 135807586 129 4 0.5334232 0.6184915 -0.0850683 -8.2364116
15366 chr3 72938383 72938510 128 4 0.9464335 0.9052487 0.0411847 8.2330861
16095 chr3 177542051 177542238 188 4 0.5962185 0.6838908 -0.0876723 -8.2259452
12014 chr2 68870123 68870193 71 4 0.9052234 0.8463731 0.0588504 8.2238674
22999 chr9 14910371 14910445 75 4 0.8438695 0.8909468 -0.0470773 -8.2224679
20264 chr6 158829586 158829684 99 4 0.5759232 0.6481398 -0.0722166 -8.2219505
2605 chr10 49997857 49997997 141 4 0.5175117 0.5992278 -0.0817161 -8.2213185
12089 chr2 77317406 77317554 149 4 0.7758259 0.7031693 0.0726566 8.2202925
7334 chr14 104766708 104766828 121 4 0.5204810 0.5863581 -0.0658771 -8.2178161
4483 chr11 117668270 117668336 67 4 0.7387075 0.6813837 0.0573238 8.2163309
1344 chr1 149784257 149784357 101 4 0.9395191 0.8981054 0.0414137 8.2075479
11390 chr19 51059755 51059828 74 4 0.7009565 0.7676660 -0.0667095 -8.2027038
10792 chr19 3665846 3665918 73 4 0.6778485 0.7495270 -0.0716785 -8.2026670
23570 chr9 97807262 97807338 77 4 0.8720412 0.8143077 0.0577335 8.2003579
9631 chr17 46059753 46059877 125 4 0.6985727 0.7639736 -0.0654009 -8.1999688
14071 chr20 62800222 62800413 192 4 0.5208431 0.6039535 -0.0831104 -8.1913130
16264 chr3 197062471 197062641 171 4 0.5402380 0.6215207 -0.0812827 -8.1894939
23575 chr9 98032093 98032177 85 4 0.5812329 0.6581075 -0.0768747 -8.1893864
22118 chr8 37403012 37403062 51 4 0.6291148 0.6966813 -0.0675665 -8.1858934
1297 chr1 145968746 145968810 65 4 0.9476279 0.9118997 0.0357282 8.1834239
2635 chr10 53981774 53981986 213 4 0.5386189 0.6369124 -0.0982935 -8.1795319
3928 chr11 59319825 59319913 89 4 0.8079171 0.7420512 0.0658660 8.1788086
12300 chr2 109636816 109636956 141 4 0.7290351 0.8054952 -0.0764601 -8.1765226
13905 chr20 50723113 50723254 142 4 0.8122919 0.7573951 0.0548969 8.1762484
21691 chr7 153446206 153446256 51 4 0.7843174 0.7257193 0.0585981 8.1749782
10795 chr19 3801922 3801972 51 4 0.8135320 0.8630308 -0.0494988 -8.1745062
8871 chr16 71928553 71928676 124 4 0.9461622 0.9724247 -0.0262625 -8.1741358
3766 chr11 36399086 36399139 54 4 0.9313957 0.8964975 0.0348982 8.1623461
16049 chr3 171793751 171793913 163 4 0.5289175 0.6169373 -0.0880198 -8.1606229
3986 chr11 64067344 64067409 66 4 0.8486803 0.7970171 0.0516631 8.1557950
9711 chr17 48255861 48255937 77 4 0.8325368 0.7631927 0.0693441 8.1346112
15295 chr3 66647876 66647946 71 4 0.8986994 0.8315859 0.0671135 8.1334196
2217 chr10 1804299 1804482 184 4 0.5142643 0.5868180 -0.0725537 -8.1216047
19807 chr6 99273830 99273939 110 4 0.9393210 0.8960916 0.0432294 8.1164008
13819 chr20 45279805 45279870 66 4 0.7913174 0.7380580 0.0532594 8.1077138
16401 chr4 10033583 10033645 63 4 0.5746968 0.6640966 -0.0893998 -8.1025946
5595 chr12 113912971 113913031 61 4 0.8262071 0.7682068 0.0580003 8.0969214
17759 chr5 31194800 31194873 74 4 0.8245511 0.8724556 -0.0479045 -8.0921648
8428 chr16 21289947 21290035 89 4 0.8842972 0.8172290 0.0670681 8.0882601
5862 chr12 132109635 132109701 67 4 0.7838971 0.7130181 0.0708790 8.0861640
14313 chr21 42541037 42541090 54 4 0.9083851 0.9464668 -0.0380817 -8.0845190
20167 chr6 146755433 146755484 52 4 0.8537915 0.8015157 0.0522759 8.0819482
24606 chrX 104154635 104154690 56 4 0.9170626 0.8642760 0.0527866 8.0770087
267 chr1 17446144 17446194 51 4 0.9432662 0.9063752 0.0368910 8.0759013
20820 chr7 45962573 45962677 105 4 0.8374627 0.7754323 0.0620304 8.0756659
14907 chr3 25263500 25263576 77 4 0.5791853 0.6573472 -0.0781619 -8.0634497
2158 chr1 244175690 244175742 53 4 0.5864759 0.6550793 -0.0686034 -8.0517748
8739 chr16 56660390 56660451 62 4 0.8645404 0.7980514 0.0664891 8.0494805
10796 chr19 3986693 3986751 59 4 0.9448859 0.9036548 0.0412311 8.0475070
6560 chr13 110789620 110789675 56 4 0.8333666 0.7680296 0.0653370 8.0465895
21677 chr7 151489488 151489649 162 4 0.5439957 0.6180171 -0.0740214 -8.0425719
22229 chr8 53339014 53339084 71 4 0.5098490 0.5910357 -0.0811866 -8.0345014
9095 chr16 89753683 89753748 66 4 0.9200130 0.9522637 -0.0322506 -8.0233338
7767 chr15 60014919 60015085 167 4 0.5263982 0.6028255 -0.0764274 -8.0210897
237 chr1 16176552 16176628 77 4 0.9602187 0.9839866 -0.0237680 -8.0128883
24160 chrX 11805989 11806058 70 4 0.8228349 0.7552708 0.0675641 8.0067511
11926 chr2 60522229 60522282 54 4 0.5558434 0.6350645 -0.0792211 -8.0037391
13771 chr20 40469814 40469870 57 4 0.9524288 0.9008643 0.0515645 8.0019216
10188 chr18 11947936 11948038 103 4 0.7682212 0.7107998 0.0574214 7.9958791
7945 chr15 72476944 72476997 54 4 0.8148994 0.8674088 -0.0525094 -7.9863288
2178 chr1 246382902 246382968 67 4 0.5630894 0.6347994 -0.0717099 -7.9711725
11667 chr2 25482809 25482979 171 4 0.8688594 0.8150692 0.0537902 7.9613673
2883 chr10 80541079 80541170 92 4 0.5079209 0.5848020 -0.0768811 -7.9562177
19495 chr6 43111240 43111307 68 4 0.8426470 0.8880928 -0.0454458 -7.9163661
11378 chr19 50651249 50651308 60 4 0.6405742 0.6987802 -0.0582060 -7.8900549
7870 chr15 68127629 68127719 91 4 0.9301715 0.8953841 0.0347874 7.8734944
9070 chr16 88170026 88170089 64 4 0.5162874 0.5841265 -0.0678391 -7.8711609
10686 chr18 77091160 77091251 92 4 0.5102384 0.5903566 -0.0801183 -7.8368396
879 chr1 67893397 67894152 756 7 0.8479713 0.8668655 -0.0188942 -7.7264341
18074 chr5 79110758 79111425 668 4 0.6077635 0.6721968 -0.0644333 -5.9280736
20118 chr6 140202506 140203171 666 5 0.8401799 0.8653739 -0.0251940 -4.9064541
6992 chr14 63513602 63514195 594 8 0.7995103 0.7811978 0.0183125 2.8611907
20502 chr7 13071723 13072314 592 4 0.8097788 0.8276757 -0.0178970 -0.1985758

DMLs:

kable(dmls) %>%
  kable_styling(bootstrap_options = "striped", full_width = F) %>%
  scroll_box(width = "100%", height = "200px")
chr pos mu1 mu2 diff diff.se stat phi1 phi2 pval fdr postprob.overThreshold
312380 chr1 20577038 0.5404625 0.9579755 -0.4175130 0.0333077 -12.535015 0.0067415 0.0067415 0.0000000 0.0000000 1.0000000
6723308 chr15 22681253 0.9779333 0.5508847 0.4270486 0.0376574 11.340359 0.0067415 0.2982368 0.0000000 0.0000000 1.0000000
8177551 chr16 73584027 0.5413766 0.9060981 -0.3647215 0.0297656 -12.253140 0.0067415 0.0067415 0.0000000 0.0000000 1.0000000
8177552 chr16 73584058 0.5413766 0.9060981 -0.3647215 0.0303462 -12.018681 0.0067415 0.0067415 0.0000000 0.0000000 1.0000000
8177553 chr16 73584069 0.5413766 0.9060981 -0.3647215 0.0297498 -12.259644 0.0067415 0.0067415 0.0000000 0.0000000 1.0000000
8177554 chr16 73584109 0.5354721 0.8878909 -0.3524188 0.0292866 -12.033456 0.0067415 0.0067415 0.0000000 0.0000000 1.0000000
8177555 chr16 73584126 0.5354721 0.8878909 -0.3524188 0.0296070 -11.903207 0.0067415 0.0067415 0.0000000 0.0000000 1.0000000
14744476 chr3 73521543 0.6207886 0.9534183 -0.3326297 0.0273828 -12.147382 0.0067415 0.0067415 0.0000000 0.0000000 1.0000000
15705174 chr4 9359173 0.6304883 0.9901312 -0.3596429 0.0307179 -11.707920 0.2037879 0.0067415 0.0000000 0.0000000 1.0000000
15705222 chr4 9363918 0.5635783 0.9822521 -0.4186738 0.0327887 -12.768851 0.4920031 0.0067415 0.0000000 0.0000000 1.0000000
15705223 chr4 9363930 0.5635783 0.9822521 -0.4186738 0.0373176 -11.219211 0.0067415 0.0009236 0.0000000 0.0000000 1.0000000
16040120 chr4 67972875 0.5609346 0.9896002 -0.4286656 0.0351706 -12.188194 0.0067415 0.0067415 0.0000000 0.0000000 1.0000000
16040121 chr4 67972901 0.5609346 0.9896002 -0.4286656 0.0339474 -12.627335 0.0067415 0.0067415 0.0000000 0.0000000 1.0000000
17016782 chr5 28864335 0.5349207 0.9800721 -0.4451513 0.0309841 -14.367107 0.0067415 0.0067415 0.0000000 0.0000000 1.0000000
17016783 chr5 28864377 0.5349207 0.9800721 -0.4451513 0.0326631 -13.628576 0.0067415 0.0067415 0.0000000 0.0000000 1.0000000
19587425 chr7 27028571 0.5805016 0.9499756 -0.3694740 0.0309058 -11.954861 0.0067415 0.0067415 0.0000000 0.0000000 1.0000000
19587426 chr7 27028574 0.5805016 0.9499756 -0.3694740 0.0302570 -12.211201 0.0067415 0.0067415 0.0000000 0.0000000 1.0000000
4501313 chr12 41739142 0.5199090 0.9126618 -0.3927528 0.0355512 -11.047521 0.0067415 0.0067415 0.0000000 0.0000000 1.0000000
5653014 chr13 71191079 0.5118941 0.9037881 -0.3918940 0.0352702 -11.111202 0.0067415 0.0067415 0.0000000 0.0000000 1.0000000
8177556 chr16 73584136 0.5354721 0.8878909 -0.3524188 0.0306380 -11.502664 0.0067415 0.0067415 0.0000000 0.0000000 1.0000000
12371634 chr2 202672283 0.5686239 0.9392053 -0.3705814 0.0330129 -11.225351 0.0067415 0.0067415 0.0000000 0.0000000 1.0000000
14744474 chr3 73521416 0.6323536 0.9636038 -0.3312502 0.0283253 -11.694517 0.0067415 0.0067415 0.0000000 0.0000000 1.0000000
19798444 chr7 54932635 0.6014882 0.9665252 -0.3650370 0.0322368 -11.323608 0.0067415 0.0067415 0.0000000 0.0000000 1.0000000
5653015 chr13 71191136 0.5118941 0.9037881 -0.3918940 0.0362005 -10.825646 0.0067415 0.0067415 0.0000000 0.0000000 1.0000000
7591091 chr16 5562189 0.5574387 0.9062097 -0.3487710 0.0309395 -11.272692 0.0067415 0.0067415 0.0000000 0.0000000 1.0000000
14744479 chr3 73521617 0.6272706 0.9584275 -0.3311569 0.0288265 -11.487930 0.0067415 0.0067415 0.0000000 0.0000000 1.0000000
19798446 chr7 54932671 0.6014882 0.9665252 -0.3650370 0.0331792 -11.001976 0.0067415 0.0067415 0.0000000 0.0000000 1.0000000
14744478 chr3 73521605 0.6272706 0.9584275 -0.3311569 0.0293937 -11.266274 0.0067415 0.0067415 0.0000000 0.0000000 1.0000000
8177557 chr16 73584186 0.5334914 0.8816945 -0.3482031 0.0316719 -10.994086 0.0067415 0.0067415 0.0000000 0.0000000 1.0000000
20995833 chr8 37005593 0.5613069 0.8947328 -0.3334258 0.0298506 -11.169810 0.0067415 0.0067415 0.0000000 0.0000000 1.0000000
14744477 chr3 73521576 0.6272706 0.9584275 -0.3311569 0.0296509 -11.168541 0.0067415 0.0067415 0.0000000 0.0000000 1.0000000
20995834 chr8 37005619 0.5613069 0.8947328 -0.3334258 0.0299532 -11.131542 0.0067415 0.0067415 0.0000000 0.0000000 1.0000000
14744475 chr3 73521447 0.6323536 0.9636038 -0.3312502 0.0299154 -11.072916 0.0067415 0.0067415 0.0000000 0.0000000 1.0000000
19798445 chr7 54932653 0.6014882 0.9665252 -0.3650370 0.0343596 -10.624033 0.0067415 0.0067415 0.0000000 0.0000000 1.0000000
14744480 chr3 73521747 0.6006706 0.9526925 -0.3520219 0.0327815 -10.738433 0.0067415 0.0067415 0.0000000 0.0000000 1.0000000
20995835 chr8 37005632 0.5616832 0.8928450 -0.3311617 0.0301708 -10.976238 0.0067415 0.0067415 0.0000000 0.0000000 1.0000000
20995830 chr8 37005555 0.5580239 0.8639182 -0.3058943 0.0272320 -11.232887 0.0067415 0.0067415 0.0000000 0.0000000 1.0000000
17402755 chr5 91443240 0.6095541 0.9553486 -0.3457945 0.0325594 -10.620420 0.0067415 0.0067415 0.0000000 0.0000000 1.0000000
20995827 chr8 37005406 0.5432692 0.8850128 -0.3417436 0.0325211 -10.508349 0.0067415 0.0067415 0.0000000 0.0000000 1.0000000
9204509 chr17 70774903 0.5219825 0.8754104 -0.3534279 0.0341340 -10.354147 0.0067415 0.0067415 0.0000000 0.0000000 1.0000000
20995831 chr8 37005558 0.5580239 0.8639182 -0.3058943 0.0277554 -11.021076 0.0067415 0.0067415 0.0000000 0.0000000 1.0000000
9580789 chr18 22236623 0.5442539 0.9384120 -0.3941580 0.0396762 -9.934382 0.0067415 0.0067415 0.0000000 0.0000000 1.0000000
9204505 chr17 70774806 0.5219825 0.8754104 -0.3534279 0.0341842 -10.338933 0.0067415 0.0067415 0.0000000 0.0000000 1.0000000
16830978 chr5 2112713 0.5200162 0.8424621 -0.3224460 0.0300498 -10.730393 0.0067415 0.0067415 0.0000000 0.0000000 1.0000000
9204507 chr17 70774869 0.5219825 0.8754104 -0.3534279 0.0342470 -10.319958 0.0067415 0.0067415 0.0000000 0.0000000 1.0000000
12545844 chr2 225803471 0.5920363 0.9421776 -0.3501413 0.0341259 -10.260290 0.0067415 0.0067415 0.0000000 0.0000000 1.0000000
9204508 chr17 70774872 0.5219825 0.8754104 -0.3534279 0.0346090 -10.212032 0.0067415 0.0067415 0.0000000 0.0000000 1.0000000
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22393351 chr9 103799933 0.5248541 0.6964982 -0.1716441 0.0434012 -3.954820 0.0067415 0.0067415 0.0000766 0.1650598 0.9506040
916953 chr1 91322473 0.5124043 0.6730930 -0.1606887 0.0367711 -4.369979 0.0067415 0.0067415 0.0000124 0.0391720 0.9505744
4971379 chr12 108866671 0.5175997 0.6875757 -0.1699760 0.0424009 -4.008787 0.0067415 0.0067415 0.0000610 0.1384423 0.9505637
19920422 chr7 71078420 0.6700064 0.8426035 -0.1725972 0.0439900 -3.923558 0.0067415 0.0067415 0.0000873 0.1824309 0.9505605
9730814 chr18 46386209 0.6782717 0.8262871 -0.1480155 0.0290959 -5.087157 0.0067415 0.0067415 0.0000004 0.0021837 0.9505539
12215259 chr2 176296138 0.5161717 0.7672318 -0.2510601 0.0915747 -2.741589 0.0067415 0.0067415 0.0061143 1.0000000 0.9505491
18280934 chr6 22149875 0.5191795 0.6861705 -0.1669910 0.0405962 -4.113460 0.0067415 0.0067415 0.0000390 0.0975094 0.9505467
14609947 chr3 54912661 0.5168666 0.6819536 -0.1650870 0.0394460 -4.185135 0.0067415 0.0067415 0.0000285 0.0761396 0.9505312
4969954 chr12 108665978 0.5212659 0.7004393 -0.1791734 0.0479836 -3.734056 0.0067415 0.0067415 0.0001884 0.3308288 0.9505296
4862957 chr12 94178452 0.5927233 0.7497179 -0.1569946 0.0345437 -4.544807 0.0067415 0.0067415 0.0000055 0.0205035 0.9505209
9697667 chr18 42477138 0.5156495 0.6773368 -0.1616873 0.0373898 -4.324375 0.0067415 0.0067415 0.0000153 0.0463533 0.9505127
7659164 chr16 12207464 0.5122514 0.6696084 -0.1573570 0.0347721 -4.525380 0.0067415 0.0067415 0.0000060 0.0220663 0.9504786
15016693 chr3 124721573 0.5454577 0.7110939 -0.1656362 0.0397917 -4.162585 0.0067415 0.0067415 0.0000315 0.0823414 0.9504770
1031646 chr1 110196047 0.5397975 0.7106614 -0.1708638 0.0429626 -3.977034 0.0067415 0.0067415 0.0000698 0.1537840 0.9504702
6554212 chr14 96562344 0.5604880 0.7296793 -0.1691913 0.0419555 -4.032634 0.0067415 0.0067415 0.0000552 0.1277105 0.9504423
12626963 chr2 235076749 0.5423015 0.7183004 -0.1759989 0.0460848 -3.819025 0.0067415 0.0067415 0.0001340 0.2543477 0.9504375
6854106 chr15 36069337 0.5060739 0.6721693 -0.1660953 0.0400804 -4.144055 0.0067415 0.0067415 0.0000341 0.0880564 0.9504332
1623956 chr1 211112959 0.5171447 0.6891375 -0.1719928 0.0436584 -3.939512 0.0067415 0.0067415 0.0000816 0.1734050 0.9504264
8074442 chr16 64931382 0.5177728 0.6895093 -0.1717365 0.0435036 -3.947637 0.0067415 0.0067415 0.0000789 0.1690444 0.9504240
21432027 chr8 104164520 0.5332074 0.6990335 -0.1658261 0.0399221 -4.153741 0.0067415 0.0067415 0.0000327 0.0849647 0.9504122
5091662 chr12 119740218 0.6878697 0.8587315 -0.1708618 0.0429777 -3.975596 0.0067415 0.0067415 0.0000702 0.1545039 0.9504063
11998961 chr2 140563267 0.5243660 0.6883043 -0.1639383 0.0387847 -4.226884 0.0067415 0.0067415 0.0000237 0.0659980 0.9503796
20417456 chr7 135822085 0.5188439 0.6902633 -0.1714194 0.0433243 -3.956655 0.0067415 0.0067415 0.0000760 0.1640774 0.9503731
7980570 chr16 52774814 0.5273906 0.6951798 -0.1677892 0.0411265 -4.079828 0.0067415 0.0067415 0.0000451 0.1094025 0.9503552
7233194 chr15 78615173 0.6852827 0.8406334 -0.1553507 0.0335847 -4.625643 0.0067415 0.0067415 0.0000037 0.0150099 0.9503333
16032466 chr4 66279813 0.5213881 0.6896727 -0.1682846 0.0414386 -4.061062 0.0067415 0.0067415 0.0000489 0.1165108 0.9503084
11545234 chr2 80679677 0.5230874 0.6903546 -0.1672672 0.0408257 -4.097105 0.0067415 0.0067415 0.0000418 0.1032408 0.9502895
2991185 chr10 127747322 0.5608338 0.7217377 -0.1609039 0.0369639 -4.352995 0.0067415 0.0067415 0.0000134 0.0417147 0.9502883
7395009 chr15 95998035 0.5172173 0.6723630 -0.1551457 0.0334704 -4.635316 0.0067415 0.0067415 0.0000036 0.0144387 0.9502824
4333329 chr12 12838483 0.7247850 0.8877023 -0.1629172 0.0381909 -4.265859 0.0067415 0.0067415 0.0000199 0.0574992 0.9502660
19232838 chr6 163158221 0.5080861 0.7576828 -0.2495968 0.0908407 -2.747633 0.0067415 0.0067415 0.0060027 1.0000000 0.9502603
6997964 chr15 53351593 0.5098398 0.6713749 -0.1615351 0.0373547 -4.324359 0.0067415 0.0067415 0.0000153 0.0463533 0.9502538
9372473 chr17 79773089 0.5668899 0.7280913 -0.1612014 0.0371526 -4.338903 0.0067415 0.0067415 0.0000143 0.0439604 0.9502517
18397935 chr6 33811854 0.5111068 0.6708875 -0.1597807 0.0362916 -4.402693 0.0067415 0.0067415 0.0000107 0.0346871 0.9502449
5037239 chr12 114072556 0.5350608 0.7197459 -0.1846851 0.0514126 -3.592211 0.0067415 0.0067415 0.0003279 0.5030693 0.9502379
19233257 chr6 163235838 0.5084534 0.6675851 -0.1591317 0.0358993 -4.432726 0.0067415 0.0067415 0.0000093 0.0310558 0.9502369
4008108 chr11 115426553 0.5326371 0.7042769 -0.1716398 0.0434940 -3.946286 0.0067415 0.0067415 0.0000794 0.1697809 0.9502331
2274629 chr10 35078956 0.5234167 0.6987486 -0.1753319 0.0457358 -3.833584 0.0067415 0.0067415 0.0001263 0.2428025 0.9502325
20835185 chr8 17270047 0.6518465 0.8060606 -0.1542141 0.0329152 -4.685194 0.0067415 0.0067415 0.0000028 0.0118152 0.9502297
6429 chr1 766285 0.5115099 0.6786283 -0.1671185 0.0407532 -4.100749 0.0067415 0.0067415 0.0000412 0.1020260 0.9502160
19904547 chr7 69329268 0.5149903 0.7663667 -0.2513764 0.0919527 -2.733756 0.0067415 0.0067415 0.0062616 1.0000000 0.9502093
14072542 chr22 45795742 0.5077418 0.6808478 -0.1731059 0.0443906 -3.899612 0.0067415 0.0067415 0.0000963 0.1971700 0.9502087
13060809 chr20 38506331 0.5770902 0.7506710 -0.1735808 0.0446855 -3.884498 0.0067415 0.0067415 0.0001025 0.2070107 0.9501836
914085 chr1 91003852 0.5947130 0.7635949 -0.1688819 0.0418322 -4.037130 0.0067415 0.0067415 0.0000541 0.1257847 0.9501824
19046481 chr6 139693248 0.7036083 0.8849166 -0.1813083 0.0493873 -3.671152 0.0067415 0.0067415 0.0002415 0.3996264 0.9501532
1691155 chr1 220826619 0.5134477 0.6710976 -0.1576499 0.0350204 -4.501661 0.0067415 0.0067415 0.0000067 0.0241242 0.9501367
22336269 chr9 97788178 0.5424785 0.7014668 -0.1589883 0.0358336 -4.436851 0.0067415 0.0067415 0.0000091 0.0306034 0.9501360
3908482 chr11 100016187 0.5187594 0.6812327 -0.1624733 0.0379516 -4.281060 0.0067415 0.0067415 0.0000186 0.0544206 0.9501313
624060 chr1 46682947 0.5250519 0.7642906 -0.2392387 0.0846099 -2.827550 0.0067415 0.0067415 0.0046906 1.0000000 0.9501131
5090343 chr12 119592109 0.8950865 0.7496602 0.1454263 0.0275993 5.269209 0.0067415 0.0067415 0.0000001 0.0009726 0.9501103
20386416 chr7 131686173 0.5804559 0.7591327 -0.1786768 0.0478019 -3.737858 0.0067415 0.0067415 0.0001856 0.3270439 0.9501070
17739384 chr5 142314181 0.5224315 0.7094798 -0.1870483 0.0528929 -3.536358 0.0067415 0.0067415 0.0004057 0.5907995 0.9500920
11914967 chr2 129402177 0.7308971 0.8924225 -0.1615254 0.0373879 -4.320260 0.0067415 0.0067415 0.0000156 0.0470923 0.9500766
14658607 chr3 61497978 0.5123202 0.6877581 -0.1754379 0.0458441 -3.826834 0.0067415 0.0067415 0.0001298 0.2480479 0.9500698
605217 chr1 45069343 0.5337099 0.7054433 -0.1717334 0.0435935 -3.939426 0.0067415 0.0067415 0.0000817 0.1734066 0.9500673
17514781 chr5 112395122 0.6142770 0.7788958 -0.1646188 0.0392772 -4.191205 0.0067415 0.0067415 0.0000277 0.0745599 0.9500356
6374045 chr14 74607040 0.5123285 0.6798164 -0.1674879 0.0410251 -4.082568 0.0067415 0.0067415 0.0000445 0.1083831 0.9500190
4028749 chr11 117544382 0.5221203 0.6866981 -0.1645778 0.0392566 -4.192364 0.0067415 0.0067415 0.0000276 0.0742298 0.9500171
13892916 chr22 33277310 0.6685415 0.8330443 -0.1645029 0.0392124 -4.195173 0.0067415 0.0067415 0.0000273 0.0734864 0.9500110

DMR calling with dmrseq

A popular R package for DMR calling is dmrseq. For more detailed description of the possibilites, see the vignette.

if(!requireNamespace("dmrseq")) {
BiocManager::install("dmrseq")
}
library(dmrseq)

register(MulticoreParam(2))

testCovariate <- "Day"
regions <- dmrseq(bs=bs_obj[240001:260000,],
                  cutoff = 0.05,
                  testCovariate=testCovariate, verbose = T)
saveRDS(regions, "dmrseq_regions.RDS")

Visualizing DMRs and candidate regions

Visualizing a region

We can visualize certain regions and/or DMRs using the Gviz package. The plotting function is included in the best_practices_WGBS.Rmd file.

dmrs <- makeGRangesFromDataFrame(dmrs, keep.extra.columns = T)
genome <- "hg19"
mart <- useMart(biomart="ENSEMBL_MART_ENSEMBL", host="feb2014.archive.ensembl.org", dataset="hsapiens_gene_ensembl")



i <- 5

#candidate_region <- GRanges(paste0(seqlevels(dmrs)[i], ":", start(dmrs)[i], "-", end(dmrs)[i]))
candidate_region <- dmrs[i]
region_plot(m = meth, region=candidate_region, mart=mart, genome=genome, groups=meth$Day, dmrs=dmrs)
#the region_plot function is available in the best_practices.Rmd file. 

Heatmap of DMRs

heatmap_data <- as.data.frame(methrix::get_region_summary(meth, makeGRangesFromDataFrame(dmrs)))
heatmap_data <- heatmap_data[complete.cases(heatmap_data),]
rownames(heatmap_data) <- paste0(heatmap_data$chr,":", heatmap_data$start, "-", heatmap_data$end)
pheatmap::pheatmap(heatmap_data[,-(1:5)], show_rownames = FALSE, annotation_col = as.data.frame(meth@colData), 
                   fontsize = 8)

Number and size of DMRs

if(!requireNamespace("plotly")) {
install.packages("plotly")
}
if(!requireNamespace("ggplot2")) {
install.packages("ggplot2")
}
if(!requireNamespace("scales")) {
install.packages("scales")
}
library(plotly)
library(ggplot2)
library(scales)
count <- c("Higher methylation in day 14" = length(dmrs[dmrs$diff.Methy<0,]),
           "Higher methylation in day 6" = length(dmrs[dmrs$diff.Methy>0,]))
length <- c("Higher methylation in day 14" = sum(width(dmrs[dmrs$diff.Methy<0,])), 
            "Higher methylation in day 6" = sum(width(dmrs[dmrs$diff.Methy<0,]))) 


data <- data.frame(Direction=c("Gain", "Loss"), Count=count, Length=length)

   g <- ggplot(data=data)+geom_col(aes(x=factor(1), y=Count, fill=Direction), position = "dodge")+theme_bw()+theme(axis.title.x = element_blank(), axis.text.x = element_blank())+scale_fill_brewer(palette = "Dark2")+ggtitle("Number of differentially methylated regions")+scale_y_continuous(labels = comma)
ggplotly(g)
   p <- ggplot(data=data)+geom_col(aes(x=factor(1), y=Length, fill=Direction), position = "dodge")+theme_bw()+theme(axis.title.x = element_blank(), axis.text.x = element_blank())+scale_fill_brewer(palette = "Dark2")+ggtitle("Length of differentially methylated regions")+scale_y_continuous(labels = comma)
ggplotly(p)

Region annotation

To describe and interpret the differentially methylated regions, we can use the ChIPseeker package. The location of the DMRs can already give us a hint the involved processes, but once we assign the regions to genes, a pathway enrichment analysis is also possible. For additional details, see the vignette of ChIPseeker:

## loading packages
if(!requireNamespace("ChIPseeker")) {
BiocManager::install("ChIPseeker")
}
if(!requireNamespace("TxDb.Hsapiens.UCSC.hg19.knownGene")) {
BiocManager::install("TxDb.Hsapiens.UCSC.hg19.knownGene")
}
if(!requireNamespace("ReactomePA")) {
BiocManager::install("ReactomePA")
}
if(!requireNamespace("clusterProfiler")) {
BiocManager::install("clusterProfiler")
}
library(ChIPseeker)
library(TxDb.Hsapiens.UCSC.hg19.knownGene)
txdb <- TxDb.Hsapiens.UCSC.hg19.knownGene
library(ReactomePA)
library(clusterProfiler)
  peakAnnoList <- lapply(list("Higher in day 14"=dmrs[dmrs$diff.Methy<0,], "Higher in day 6" = dmrs[dmrs$diff.Methy>0,]), annotatePeak, TxDb=txdb, tssRegion=c(-3000, 3000), verbose=FALSE)

plotAnnoBar(peakAnnoList)

plotDistToTSS(peakAnnoList)

genes <- lapply(list("Higher in day 14"=dmrs[dmrs$diff.Methy<0,], "Higher in day 6" = dmrs[1:3,]), seq2gene, tssRegion = c(-1000, 1000), flankDistance = 3000, TxDb=txdb)
##   403 genes were dropped because they have exons located on both strands
##   of the same reference sequence or on more than one reference sequence,
##   so cannot be represented by a single genomic range.
##   Use 'single.strand.genes.only=FALSE' to get all the genes in a
##   GRangesList object, or use suppressMessages() to suppress this message.
names(genes) <- gsub(" ", "\n", names(genes))
comppthw <- compareCluster(geneCluster   = genes,
                         fun           = "enrichPathway",
                         pvalueCutoff  = 0.05,
                         pAdjustMethod = "BH")
dotplot(comppthw, font.size = 8)

gc()
##              used    (Mb) gc trigger    (Mb)   max used    (Mb)
## Ncells   13096625   699.5   38982997  2082.0   48728746  2602.4
## Vcells 2214549494 16895.7 3526654847 26906.3 3526648952 26906.2

Working with HDF5-based object

read_bedgraph function

For memory efficient read in, one can use an HDF5 based methrix object. Only one bedgraph file is in the memory at the same time, while the resulting object won’t be stored in the memory, but on-disk. Additional arguments to use HDF5: * Set h5=TRUE * Set vect=FALSE * h5_dir –> a directory to save the final object. It is possible to save the object later. It increases the processing time significantly.
* h5temp –> a temporary directory to use during data processing. Set this if for example the default temporary location doesn’t have enough free space to store the temporary data.

meth <- methrix::read_bedgraphs(
  files = bed_files,
  ref_cpgs = hg19_cpgs,
  chr_idx = 1,
  start_idx = 2,
  M_idx = 5,
  U_idx = 6,
  stranded = FALSE,
  zero_based = TRUE, 
  collapse_strands = FALSE,  
  coldata = sample_anno,
  vect = FALSE,
  h5 = TRUE)
## ----------------------------
## -Preset:        Custom
## --Missing beta and coverage info. Estimating them from M and U values
## -CpGs raw:      29,891,155 (total reference CpGs)
## -CpGs retained: 28,217,448(reference CpGs from contigs of interest)
## ----------------------------
## -Processing:    GSM3039348_BiSeq_WT_D14_R1.bed.gz
## --CpGs missing: 291,768 (from known reference CpGs)
## -Processing:    GSM3039349_BiSeq_WT_D14_R2.bed.gz
## --CpGs missing: 213,929 (from known reference CpGs)
## -Processing:    GSM3039350_BiSeq_WT_D14_R3.bed.gz
## --CpGs missing: 391,604 (from known reference CpGs)
## -Processing:    GSM3039352_BiSeq_WT_D6_R1.bed.gz
## --CpGs missing: 288,499 (from known reference CpGs)
## -Processing:    GSM3039353_BiSeq_WT_D6_R2.bed.gz
## --CpGs missing: 225,954 (from known reference CpGs)
## -Processing:    GSM3039354_BiSeq_WT_D6_R3.bed.gz
## --CpGs missing: 392,670 (from known reference CpGs)
## -Finished in:  00:06:22 elapsed (00:06:37 cpu)

All methrix functions work with HDF5-based objects as well, there is no difference in using different functions.

meth <- methrix::remove_uncovered(meth)
## -Removed 113,365 [0.4%] uncovered loci of 28,217,448 sites
## -Finished in:  8.660s elapsed (7.060s cpu)

It is also possible to transform non-HDF5-based objects to HDF5-based ones and back.

m <- convert_HDF5_methrix(m=meth)
m2 <- convert_methrix(m=m)

Saving and loading

Saving and loading of an HDF5-based object is not possible using the standard save or saveRDS functions. methrix offers easy to use saving and loading tools, which are essentially wrappers around the saveHDF5SummarizedExperiment and loadHDF5SummarizedExperiment functions.

target_dir = paste0( getwd(), '/temp/')
save_HDF5_methrix(meth, dir = target_dir, replace = TRUE)

meth <- load_HDF5_methrix(dir = target_dir)

Working with large number of samples

The primary goal of methrix is to allow users to handle the whole-genome methylation data. The functions are optimized to keep the speed high and the memory need low. However, additional efforts were taken to allow methrix to handle large number of samples (even >100) in the samples in the same, efficient way. Therefore, many functions implement the argument n_chunks to split these datasets into digestible chunks and n_cores to parallelize the processing of these chunks. Functions currently supporting the arguments n_chunks and n_cores: coverage_filter get_region_summary remove_snps mask_methrix only support the n_cores argument.

The multicore option is not available on Windows.

if (grepl("Windows", Sys.getenv("OS"))){
res <- get_region_summary(meth, regions=dmrs[1:5], n_chunks = 2, n_cores = 1)} else {
  res <- get_region_summary(meth, regions=dmrs[1:5], n_chunks = 2, n_cores = 2)
}
## -Checking for overlaps..
## -Summarizing by average
## -Done! Finished in:35.9s elapsed (34.6s cpu)
res
##      chr     start       end n_overlap_CpGs rid WT_D14_R1 WT_D14_R2 WT_D14_R3
## 1:  chr5  92930350  92933392            129   1 0.8811686 0.8173702 0.9137123
## 2: chr19   1466497   1467764            133   2 0.7838913 0.7717582 0.8255125
## 3:  chr5 172110452 172111266            135   3 0.7495186 0.6830135 0.8394217
## 4:  chr9  67340092  67341092            129   4 0.7024922 0.6966098 0.7828817
## 5:  chr9 140917238 140918281            133   5 0.7666572 0.8951001 0.9316220
##     WT_D6_R1  WT_D6_R2  WT_D6_R3
## 1: 0.9474634 0.9223326 0.9368258
## 2: 0.8456527 0.8533729 0.9048029
## 3: 0.8279274 0.7470575 0.9463241
## 4: 0.7753189 0.7799586 0.8612769
## 5: 0.9360165 0.8885757 0.9528225

Integration with other tools

methrix has multiple options to export data for different purposes. The write_bedgraph function supports a bedgraph format, that can be directly uploaded to UCSC. The multibed option generates a file with all samples as columns. The metilene option ensures that this file is in the correct format for metilene, a command line DMR caller.

# UCSC browser
write_bedgraphs(m = meth, output_dir = ".")

# Metilene
write_bedgraphs(m = meth, output_dir = ".", rm_NA = FALSE, metilene = TRUE,multiBed = "metline_ip", phenoCol = "Day")

We also support bigWig format.

write_bigwigs(meth, output_dir = ".", samp_names = NULL)